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[山地山毛榉林地面植被中不同生命形式的二氧化碳交换对光照和温度的影响]

[The effect of light and temperature of the CO exchange of different life forms in the ground vegetation of a montane beech forest].

作者信息

Schulze E-D

机构信息

Botanisches Institut II der Universität Würzburg, Würzburg, Deutschland.

出版信息

Oecologia. 1972 Sep;9(3):235-258. doi: 10.1007/BF00345234.

Abstract

In a montane beech (Fagus sylvatica) forest the influence of the climatic factors, light and temperature, on net photosynthesis and on the CO balance of the ground vegetation was investigated. The total turnover of carbon was calculated. Species studied included: Athyrium filix-femina, Oxalis acetosella, Luzula luzuloides, Deschampsia flexuosa and young plants of Fagus sylvatica. 1. The light compensation point in all spp. is between 300 and 500 lux except for D. flexuosa where it is 2 klx. Light saturation is attained at 2-3 klx for A. filix-femina, at 5-6 klx for O. acetosella, and at 6-7 klx for L. luzuloides and F. sylvatica. The net photosynthesis of D. flexuosa increases linearly upto 12 klx. This plant, therefore, is more closely related to plants with high light requirements than all the other species under experiment. 2. The maximum rates of net photosynthesis in O. acetosella and A. filix-femina are higher than in all the other plants, independent of the reference system. Per unit dry weight they even attain rates of CO uptake (22-27 mg CO/gdw·h) known from herbs under the much better light conditions of an open habitat. F. sylvatica and L. luzuloides exhibit per unit dry weight only 30% of this rate and D. flexuosa 25%. On a leaf surface area and chlorophyll content basis differences are smaller: F. sylvatics attains 75%, L. luzuloides reaches 50% and D. flexuosa only 30% of the maximal rates of net photosynthesis of O. acetosella and A. filix-femina. The higher CO uptake of O. acetosella and A. filix-femina points to a better adaptation of their photosynthetic apparatus in comparison to all the other species of the same habitat. 3. At light saturation the temperature optimum of A. filix-femina and O. acetosella covers a smaller range at lower temperatures than was found in the other species. These attain almost maximal rates of net photosynthesis over the whole range of temperatures of their natural habitat. At decreasing light intensities the temperature optimum of O. acetosella changes from 13-18° C at 8-12 klx to a range of even lower temperatures (9-12° C at 1 klx). 4. The respiration of the rhizome and the roots of O. acetosella is per unit dry weight 40% of the dark respiration rate in the above ground material. 5. The daily gain of net photosynthesis per unit dry weight of O. acetosella and A. filix-femina is 4 times as high as in L. luzuloides and in F. sylvatica and 7 times as high as in D. flexuosa. Per unit of surface area and chlorophyll content differences are smaller. The sequence in all cases remains the same. During the night D. flexuosa has the highest relative respiratory loss. Its CO gain over 24 hours is very small. 6. The importance of sun flecks on the CO balance is small in all species except D. flexuosa. More important is the mean light intensity and the rate of net photosynthesis which is attained under these conditions. The amount of CO photosynthetically bound in sun flecks is 6% of the daily balance in A. filix-femina, 16-19% of the daily balance in O. acetosella, L. luzuloides and F. sylvatica, and 27% of the daily balance in D. flexuosa. The existence of D. flexuosa is dependent on the occurrence of sun flecks on the forest floor. The ecological significance of the relative light intensity in the mosaic-like distribution of plants on the forest floor is discussed. 7. The varying success in adaptation to the conditions of the habitat becomes even more evident when compared with the primary production of the beech crown. The daily gain of net photosynthesis of O. acetosella and A. filix-femina per unit dry weight is much larger than in either the sun or shade leaves in the canopy of the same stand. Per unit surface area of the leaves they attain 18-20%, per unit chlorophyll content 32% (L. luzuloides and the young plants of F. sylvatica 16-27%, D. flexuosa 4%) of the gain of net photosynthesis in the beech sun leaves. 8. A comparison with a model of primary production (maximal rates of net photosynthesis under experimentally optimal conditions over the whole day = 100%) shows what effect the different climatic factors of the natural habitat have in limiting the CO balance, and to what extend the actual CO gain reaches the physiological optinum. On the forest floor the rate of net photosynthesis is reduced primarily through the intense shade of the beech canopy and by dawn and dusk (reduction of the maximal CO gain in O. acetosella and in A. filix-femina ca. 50%, in L. luzuloides and F. sylvatica ca. 60% and in D. flexuosa ca. 86%). The effect of additional clouds is smaller (reduction of the maximal CO gain in all species 4-6%, maximal 19%). The effect of temperature is very small for L. luzuloides, F. sylvatica and D. flexuosa. Corresponding to the low temperature optimum the influence of the prevailing temperatures is much higher in O. acetosella and A. filix-femina (4-6% reduction of the maximal CO gain through temperatures above optimum). 9. For an assessment of the competition potential of O. acetosella compared to L. luzuloides and F. sylvatica for the vegetation period of April to August, the carbon balances of the whole plants were estimated. These are compared with the dry weight increase. Among the three life forms of the rhizome geophyte (Oxalis), the hemicryptophyte (Luzula) and the phanerophyte (Fagus) there are striking differences in the use and in the distribution of the CO gain. L. luzuloides invests 65% of the net photosynthetic gain as dry weight increment (O. acetosella only 44%, F. sylvatica 40%). Moreover, the growth of L. luzuloides takes place primarily above ground with the establishment of new leaves. The relative proportion of the growth above ground to the total dry weight increment is for L. luzuloides 63%, as against 57% in F. sylvatica and only 42% in O. acetosella. In respect to the total carbon balance the better use of the CO gain gives L. luzuloides in this habitat a higher competition potential than the photosynthetically more active O. acetosella. The constitutional differences in the photosynthetic activity are compensated for by the distribution of the assimilates to shoot and root.

摘要

在一片山地山毛榉(欧洲山毛榉)森林中,研究了气候因素(光照和温度)对地面植被净光合作用及碳平衡的影响。计算了碳的总周转量。所研究的物种包括:蹄盖蕨、酢浆草、亮叶地杨梅、曲芒发草以及欧洲山毛榉的幼苗。1. 除曲芒发草的光补偿点为2千勒克斯外,所有物种的光补偿点在300至500勒克斯之间。蹄盖蕨在2至3千勒克斯达到光饱和,酢浆草在5至6千勒克斯达到光饱和,亮叶地杨梅和欧洲山毛榉在6至7千勒克斯达到光饱和。曲芒发草的净光合作用在12千勒克斯之前呈线性增加。因此,与所有其他实验物种相比,该植物与高光需求植物的关系更为密切。2. 无论参照体系如何,酢浆草和蹄盖蕨的净光合作用最大速率均高于所有其他植物。按单位干重计算,它们甚至达到了开阔生境中草本植物在更好光照条件下已知的二氧化碳吸收速率(22 - 27毫克二氧化碳/克干重·小时)。欧洲山毛榉和亮叶地杨梅按单位干重计算仅为该速率的30%,曲芒发草为25%。以叶表面积和叶绿素含量为基础,差异较小:欧洲山毛榉达到酢浆草和蹄盖蕨净光合作用最大速率的75%,亮叶地杨梅达到50%,曲芒发草仅为30%。酢浆草和蹄盖蕨较高的二氧化碳吸收表明,与同一栖息地的所有其他物种相比,它们的光合器官适应性更好。3. 在光饱和时,蹄盖蕨和酢浆草的最适温度在较低温度下的范围比其他物种小。在其自然栖息地的整个温度范围内,其他物种几乎能达到净光合作用的最大速率。随着光照强度降低,酢浆草的最适温度从8 - 12千勒克斯时的13 - 18℃变为1千勒克斯时更低的温度范围(9 - 12℃)。4. 酢浆草根茎和根的呼吸按单位干重计算是地上部分暗呼吸速率的40%。5. 酢浆草和蹄盖蕨单位干重的净光合作用日增量是亮叶地杨梅和欧洲山毛榉的4倍,是曲芒发草的7倍。以单位表面积和叶绿素含量计算,差异较小。所有情况下的顺序保持不变。夜间,曲芒发草的相对呼吸损失最高。其24小时内的二氧化碳增量非常小。6. 除曲芒发草外,所有物种中光斑对碳平衡的重要性较小。更重要的是平均光照强度以及在这些条件下达到的净光合作用速率。光斑中光合固定的二氧化碳量在蹄盖蕨中占日平衡的6%,在酢浆草、亮叶地杨梅和欧洲山毛榉中占日平衡的16 - 19%,在曲芒发草中占日平衡的27%。曲芒发草的生存依赖于森林地面光斑的出现。讨论了森林地面植物镶嵌分布中相对光照强度的生态意义。7. 与山毛榉树冠的初级生产力相比,适应栖息地条件的不同成功程度更加明显。酢浆草和蹄盖蕨单位干重的净光合作用日增量比同一林分树冠中的阳叶或阴叶都大得多。以叶单位表面积计算,它们达到山毛榉阳叶净光合作用增量的18 - (此处原文似乎不完整)2%,以单位叶绿素含量计算为32%(亮叶地杨梅和欧洲山毛榉幼苗为16 - 27%,曲芒发草为4%)。8. 与初级生产模型(全天实验最佳条件下的净光合作用最大速率 = 100%)比较表明,自然栖息地的不同气候因素对碳平衡的限制作用以及实际二氧化碳增量达到生理最佳值的程度。在森林地面,净光合作用速率主要因山毛榉树冠的强烈遮荫以及黎明和黄昏而降低(酢浆草和蹄盖蕨的最大二氧化碳增量降低约50%,亮叶地杨梅和欧洲山毛榉降低约60%,曲芒发草降低约86%)。额外云层的影响较小(所有物种的最大二氧化碳增量降低4 - 6%,最大为19%)。温度对亮叶地杨梅、欧洲山毛榉和曲芒发草的影响非常小。与低温最适值相对应,酢浆草和蹄盖蕨中盛行温度的影响更高(温度高于最适值时,最大二氧化碳增量降低4 - 6%)。9. 为评估4月至8月植被生长期间酢浆草与亮叶地杨梅和欧洲山毛榉相比的竞争潜力,估算了整株植物的碳平衡。并将其与干重增加进行比较。在根茎类地生植物(酢浆草)、半隐生植物(亮叶地杨梅)和显花植物(欧洲山毛榉)这三种生活型中,二氧化碳增量的利用和分配存在显著差异。亮叶地杨梅将净光合增量的65%用于干重增加(酢浆草仅为44%,欧洲山毛榉为40%)。此外,亮叶地杨梅的生长主要发生在地上,通过新叶的形成。地上生长占总干重增量的相对比例,亮叶地杨梅为63%,欧洲山毛榉为57%,酢浆草仅为42%。就总碳平衡而言,亮叶地杨梅在该栖息地对二氧化碳增量的更好利用使其比光合活性更高的酢浆草具有更高的竞争潜力。光合活性的结构差异通过同化物在地上和地下部分的分配得到补偿。

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